However, the Mirror lost readers on PC and in print, leaving its combined readership down by 1%.

At the other end of the market the momentum is with The Times, which slightly softened its hard paywall in July 2016 to allow non-subscribers to read two stories per week in return for registering.

This helped boost its combined readership by 16% to 5,790,000.

However, those extra 780,000 Times readers are hard to square with the ten million readers lost by three rivals: The Guardian, The Independent and The Telegraph.

The Telegraph lost nearly five million, 19% of its combined readership, and the blame was down to its PC and mobile platforms.

In November last year, too late to have much effect on the figures, it opted to switch its metered paywall for a hybrid free and subscription content approach.

The Guardian also saw its stable print readership undermined by its digital performance despite its free access website, and lost 8 per cent of its combined readership.

The Independent also lost digital readers even though it dropped its print edition in March 2016. Readership fell on PC by 17% to 4,376,000 and on mobile by 2% to 14,405,000.

The Pamco figures also cover a number of magazines, with the medium experiencing a similar move to mobile but a slight decline in combined readership to 37,691,000.

A News UK spokeswoman said: "We're immensely proud of the total brand reach of The Sun, The Sun on Sunday and thesun.co.uk. The digital team under Keith Poole have grown The Sun website from a standing start to a scaled audience in the UK and there aren't many sites that have the skill set and brand recognition to be able to do that.

"Nearly half of all of our readers choose to read the print product on a monthly basis, which means that The Sun remains the most widely-read newspaper in the UK."